The Giant Extra-Floral Nectaries of Carnivorous Heliamphora Folliculata: Architecture and Ultrastructure

Total Page:16

File Type:pdf, Size:1020Kb

The Giant Extra-Floral Nectaries of Carnivorous Heliamphora Folliculata: Architecture and Ultrastructure ACTA BIOLOGICA CRACOVIENSIA Series Botanica 49/2: 91–104, 2007 THE GIANT EXTRA-FLORAL NECTARIES OF CARNIVOROUS HELIAMPHORA FOLLICULATA: ARCHITECTURE AND ULTRASTRUCTURE BARTOSZ J. PŁACHNO1*, PIOTR ŚWIĄTEK2, AND ANDREAS WISTUBA3 1Department of Plant Cytology and Embryology, Jagiellonian University, ul. Grodzka 52, 31–044 Cracow, Poland, 2Department of Animal Histology and Embryology, University of Silesia, ul. Bankowa 9, 40–007 Katowice, Poland 3Maudauer Ring 227, 68259 Mannheim, Germany Received September 15, 2007; revision accepted December 7, 2007 Extra-floral nectaries commonly occur in carnivorous plants, forming pitfall traps to attract nectar-feeding insects. Although they are not connected with pollination, extra-floral nectaries promote the reproductive func- tions of carnivorous plants by increasing the supply of animal-sourced nutrients and thereby increasing the plant's vigor. Our main purpose here was to study the functional ultrastructure of the giant nectaries in Heliamphora, focusing on nectar production and secretion. We wanted to determine whether specialization of the shape and struc- ture of Heliamphora nectar spoons has an influence on nectary structure. Heliamphora folliculata, with its unique nectar storage chamber, may also have specialized giant nectaries differing from other species in the genus. In Heliamphora folliculata the largest nectaries occur in a nectar storage chamber. Regardless of their size, the nec- taries have similar ultrastructure. Key features of their cells are ER-sheathed leucoplasts and vacuoles with large osmiophilic phenolic inclusions. The former is characteristic for cells producing monoterpenes; indeed, the giant nectaries produce volatile compounds and may have a function similar to osmophores. Nectary cells are isolated from ordinary parenchyma cells by cutinized walls lacking plasmodesmata (endodermis). Symplastic transport is possible only between nectary cells and special parenchyma cells that have wall thickenings. Between them are many plasmodesmata; thus the nectary is a symplastic and apoplastic field. These specialized parenchyma cells are similar to the flange cells described in parasitic plants. Why has a special spoon with a nectar chamber evolved in Heliamphora folliculata? One answer given is that it protects nectar against being washed away by frequent rain- falls so that the plant produces less nectar and saves energy. Also, when nectar is not easily accessible the insects have to spend more time near the trap entrance to look for it, and they are more likely to be trapped. Regardless of the shape and structure of Heliamphora nectar spoons (pitcher appendages), giant nectaries apparently have the same architecture throughout the genus. So far as is known, pollinator-prey conflict does not exist in Heliamphora; nectaries in this genus are formed only for nectar-feeding prey. Key words: Heliamphora, Sarraceniaceae, extra-floral nectaries, nectar, carnivorous plants, ultra- structure, symplastic field, osmophores, leucoplasts, carnivorous syndrome, tepui. INTRODUCTION 2006). Like other pitcher plants (Cephalotus, Nepenthes, Darlingtonia, Sarracenia), Heliamphora The family Sarraceniaceae, with three genera possesses extra-floral nectaries on the pitcher trap (Darlingtonia, Heliamphora, Sarracenia), groups surface. Nectar-feeding insects are attracted and carnivorous plants from the New World forming trapped (Jaffe et al., 1995). Their carcasses are a mainly pitfall traps (excepting Sarracenia psittacina source of nutrients for the plants (Lloyd, 1942; with a lobster-type trap; Studnièka, 1984). According Juniper et al., 1989; Jaffe et al., 1992). The anatomy to recent molecular studies, the genus Sarracenia is of nectaries of a few species of Heliamphora were first sister to Heliamphora, and this pair is sister to the described by Kraft (1896, after Lloyd, 1942) and later genus Darlingtonia (Bayer et al., 1996; Neyland and in detail by Lloyd (1942). Nectary morphology has Merchant, 2006). The genus Heliamphora comprises been studied by SEM in only two Heliamphora about 13 species and is restricted to tepuis and species (H. heterodoxa, H. nutans), by Adams and plateaus of Venezuela, Brazil and Guyana (Rice, Smith (1977) and Juniper et al. (1989); they report- *e-mail: [email protected] PL ISSN 0001-5296 © Polish Academy of Sciences, Cracow 2007 92 Płachno et al. Fig. 1. (a) Heliamphora folliculata in its natural habitat in southern Venezuela, (b) Upper part of pitcher with nectar spoon (nectar chamber damaged), southern Venezuela, (c) Upper part of pitcher of juvenile plant, lacking the spoon; note glabrous triangular part with numerous nectaries, (d) Part of leaf of juvenile plant; on the external wings are long, straight, nonsecretory hairs together with numerous glands, (e,f) Small nectaries among downward-directed, unicellu- lar, nonsecretory epidermal hairs, in adult plant. n – nectary. Giant nectaries of Heliamphora 93 Fig. 2. (a) Section through nectar spoon. H – nectar chamber; CH – nectar channel; B – base, (b) Numerous small nec- taries on glabrous spoon base, (c) Small nectary with four apical cells and remnants of secretion, (d) Part of section through nectar chamber; giant nectaries visible (*). Note strong fluorescence of cutinized walls. Bar = 300 μm, (e) Section through nectar chamber; numerous giant nectaries visible. ed two kinds of nectaries in Heliamphora: large ones Sarraceniaceae are restricted to Heliamphora. There restricted to the pitcher appendage (spoon), and have been no ultrastructure studies of nectaries in small ones occurring on the outer and partly on the this genus. Vogel (1998) pointed to the need for work inner pitcher surface. The small nectaries have a on the detailed ultrastructure of the nectaries of other basic construction very similar to that of the extra-flo- members of Sarraceniaceae. To date, the only group ral nectaries of Sarracenia, "Sarracenia-type" (Lloyd, of carnivorous plants to be studied in detail in terms 1942; Vogel, 1998). The giant extra-floral nectaries in of nectary ultrastructure and nectar production is 94 Płachno et al. Nepenthes (Vassilyev, 1977; Juniper et al, 1989; SCANNING ELECTRON MICROSCOPY Merbach et al., 2001). The procedures for preparing samples for conven- Heliamphora folliculata Wistuba, Harbarth & tional SEM were as described earlier (Płachno et al., Carow was described only recently, in 2001, from the 2005a,b). Briefly, traps were hand-sectioned with a Los Testigos table mountains in southern Venezuela. razor blade and fixed as for TEM or in 70% ethanol This endemic species is unique not only among with 1% glycerine. The material was later dehydrat- Heliamphora species but also in the Sarraceniaceae ed in an ethanol and acetone series, and critical- family, in that its pitcher appendage (spoon) forms a point dried using liquid CO . The dried tissues were chamber to store nectar (Fig. 1a,b; Wistuba et al., 2 sputter-coated with gold and viewed in a HITACHI 2001). The structure and shape of pitcher appendages S–4700 microscope (Scanning Microscopy with nectaries is one of the most valuable characters Laboratory of Biological and Geological Sciences, for interpreting Heliamphora relationships (Wistuba Jagiellonian University). To study pit distribution et al., 2001, 2002 and Carow et al., 2005). and cell wall architecture, we extracted cytoplasm. One of our aims here is to study nectar cham- Nectar spoons were hand-sectioned and fresh tissue ber anatomy and the distribution of nectaries in was placed in commercial bleach (sodium hypochlo- Heliamphora folliculata. The main purpose is to rite) diluted 1:10 in water. After 2 h the material was examine the ultrastructure of the giant nectaries of washed in water and fixed in 2.5% glutaraldehyde in Heliamphora, focusing on nectar secretion. Another cacodylate buffer. Later procedures for preparing important aim was to determine whether specializa- samples for SEM were as above. tion of the shape and structure of Heliamphora nec- tar spoons has an influence on nectary structure. Perhaps Heliamphora folliculata, with its unique HISTOCHEMISTRY nectar storage chamber, also has specialized giant Autofluorescence (chlorophyll a, cutinized walls) nectaries differing from other species in the genus. observations of fresh sectioned tissue were made in an epifluorescence microscope (Nikon Eclipse E 400 with UV–2A and B–2A filters). For some samples MATERIALS AND METHODS DAPI was added to label DNA. Documentation was PLANT MATERIAL made on Sensia 200 and 100 slide film. Juvenile and adult plants of Heliamphora folliculata, originated from seeds collected by Andreas Wistuba RESULTS during a field trip in 2001 (Wistuba et al., 2001) from Aparaman Tepui and Murosipan Tepui, were exam- SPOON ANATOMY AND NECTARY DISTRIBUTION ined. Additionally, Heliamphora folliculata plants Leaves of juvenile plants lack the spoon; only the from the Murosipan Tepui and plants of Heliamphora apex of the pitcher is slightly curved and forms a minor (Auyan Tepui) and Heliamphora heterodoxa small appendix at the tip (Fig. 1c). There are only were obtained from the collection of Kamil Pásek of small nectaries scattered on the glabrous inner sur- Dobroslavice, Czech Republic. face of the apex, the glabrous triangular part of the upper inner pitcher wall, and the glabrous pitcher margins. As in mature traps, nectaries occur also on LIGHT AND TRANSMISSION ELECTRON MICROSCOPY the inner pitcher wall among downward-directed, uni- The procedures for preparing samples for TEM
Recommended publications
  • Carnivorous Plant Newsletter V42 N3 September 2013
    Technical Refereed Contribution Phylogeny and biogeography of the Sarraceniaceae JOHN BRITTNACHER • Ashland, Oregon • USA • [email protected] Keywords: History: Sarraceniaceae evolution The carnivorous plant family Sarraceniaceae in the order Ericales consists of three genera: Dar- lingtonia, Heliamphora, and Sarracenia. Darlingtonia is represented by one species that is found in northern California and western Oregon. The genus Heliamphora currently has 23 recognized species all of which are native to the Guiana Highlands primarily in Venezuela with some spillover across the borders into Brazil and Guyana. Sarracenia has 15 species and subspecies, all but one of which are located in the southeastern USA. The range of Sarracenia purpurea extends into the northern USA and Canada. Closely related families in the plant order Ericales include the Roridu- laceae consisting of two sticky-leaved carnivorous plant species, Actinidiaceae, the Chinese goose- berry family, Cyrillaceae, which includes the common wetland plant Cyrilla racemiflora, and the family Clethraceae, which also has wetland plants including Clethra alnifolia. The rather charismatic plants of the Sarraceniaceae have drawn attention since the mid 19th century from botanists trying to understand how they came into being, how the genera are related to each other, and how they came to have such disjunct distributions. Before the advent of DNA sequencing it was very difficult to determine their relationships. Macfarlane (1889, 1893) proposed a phylogeny of the Sarraceniaceae based on his judgment of the overlap in features of the adult pitchers and his assumption that Nepenthes is a member of the family (Fig. 1a). He based his phy- logeny on the idea that the pitchers are produced from the fusion of two to five leaflets.
    [Show full text]
  • Insectivorous Plants”, He Showed That They Had Adaptations to Capture and Digest Animals
    the Strange, the Ugly, and the Bizarre . carnivores, parasites, and mycotrophs . Plant Oddities - Carnivores, Parasites & Mycotrophs Of all the plants, the most bizarre, the least understood, but yet the most interesting are those plants that have unusual modes of nutrient uptake. Carnivore: Nepenthes Plant Oddities - Carnivores, Parasites & Mycotrophs Of all the plants, the most bizarre, the least understood, but yet the most interesting are those plants that have unusual modes of nutrient uptake. Parasite: Rafflesia Plant Oddities - Carnivores, Parasites & Mycotrophs Of all the plants, the most bizarre, the least understood, but yet the most interesting are those plants that have unusual modes of nutrient uptake. Things to focus on for this topic! 1. What are these three types of plants 2. How do they live - selection 3. Systematic distribution in general 4. Systematic challenges or issues 5. Evolutionary pathways - how did they get to what they are Mycotroph: Monotropa Plant Oddities - The Problems Three factors for systematic confusion and controversy 1. the specialized roles often involve reductions or elaborations in both vegetative and floral features — DNA also is reduced or has extremely high rates of change for example – the parasitic Rafflesia Plant Oddities - The Problems Three factors for systematic confusion and controversy 2. their connections to other plants or fungi, or trapping of animals, make these odd plants prone to horizontal gene transfer for example – the parasitic Mitrastema [work by former UW student Tom Kleist]
    [Show full text]
  • The Cost of Carnivory for Darlingtonia Californica (Sarraceniaceae): Evidence from Relationships Among Leaf Traits1
    American Journal of Botany 92(7): 1085±1093. 2005. THE COST OF CARNIVORY FOR DARLINGTONIA CALIFORNICA (SARRACENIACEAE): EVIDENCE FROM RELATIONSHIPS AMONG LEAF TRAITS1 AARON M. ELLISON2,4 AND ELIZABETH J. FARNSWORTH3 2Harvard University, Harvard Forest, P.O. Box 68, Petersham, Massachusetts 01366 USA; and 3New England Wild Flower Society, 180 Hemenway Road, Framingham, Massachusetts 01701 USA Scaling relationships among photosynthetic rate, foliar nutrient concentration, and leaf mass per unit area (LMA) have been observed for a broad range of plants. Leaf traits of the carnivorous pitcher plant Darlingtonia californica, endemic to southern Oregon and northern California, USA, differ substantially from the predictions of these general scaling relationships; net photosynthetic rates of Darlingtonia are much lower than predicted by general scaling relationships given observed foliar nitrogen (N) and phosphorus (P) concentrations and LMA. At ®ve sites in the center of its range, leaf traits of Darlingtonia were strongly correlated with elevation and differed with soil calcium availability and bedrock type. The mean foliar N : P of 25.2 6 15.4 of Darlingtonia suggested that these plants were P-limited, although N concentration in the substrate also was extremely low and prey capture was uncommon. Foliar N : P stoichiometry and the observed deviation of Darlingtonia leaf traits from predictions of general scaling relationships permit an initial assessment of the ``cost of carnivory'' in this species. Carnivory in plants is thought to have evolved in response to N limitation, but for Darlingtonia, carnivory is an evolutionary last resort when both N and P are severely limiting and photosynthesis is greatly reduced. Key words: carnivorous plants; Darlingtonia californica; fens; leaf mass area; leaf traits; photosynthesis; nitrogen; serpentine.
    [Show full text]
  • Redalyc.Overcoming DNA Extraction Problems from Carnivorous Plants
    Anales del Jardín Botánico de Madrid ISSN: 0211-1322 [email protected] Consejo Superior de Investigaciones Científicas España Fleischmann, Andreas; Heubl, Günther Overcoming DNA extraction problems from carnivorous plants Anales del Jardín Botánico de Madrid, vol. 66, núm. 2, julio-diciembre, 2009, pp. 209-215 Consejo Superior de Investigaciones Científicas Madrid, España Available in: http://www.redalyc.org/articulo.oa?id=55612913003 How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative Anales del Jardín Botánico de Madrid Vol. 66(2): 209-215 julio-diciembre 2009 ISSN: 0211-1322 doi: 10.3989/ajbm.2198 Overcoming DNA extraction problems from carnivorous plants by Andreas Fleischmann & Günther Heubl LMU Munich, Systematic Botany and Mycology, Menzinger Strasse 67, D-80638 Munich, Germany [email protected] Abstract Resumen Fleischmann, A. & Heubl, G. 2009. Overcoming DNA extraction Fleischmann, A. & Heubl, G. 2009. Superando problemas de ex- problems from carnivorous plants. Anales Jard. Bot. Madrid tracción de ADN de plantas carnívoras. Anales Jard. Bot. Madrid 66(2): 209-215. 66(2): 209-215 (en inglés). We tested previously published protocols for DNA isolation from Probamos algunos protocolos publicados previamente para el plants with high contents of polyphenols and polysaccharides aislamiento del ADN de plantas con alto contenido de polifeno- for several taxa of carnivorous plants. However, we did not get les y polisacáridos para varios táxones de plantas carnívoras. Sin satisfying results with fresh or silica dried leaf tissue obtained embargo, no conseguimos muy buenos resultados ni con tejidos from field collected or greenhouse grown plants, nor from de hojas frescas, ni con tejidos de hojas secadas en gel de sílice herbarium specimens.
    [Show full text]
  • Biljke Mesožderke Carnivorous Plants
    View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Croatian Digital Thesis Repository SVEUČILIŠTE U ZAGREBU PRIRODOSLOVNO-MATEMATIČKI FAKULTET BIOLOŠKI ODSJEK BILJKE MESOŽDERKE CARNIVOROUS PLANTS Tihana Jelačić Preddiplomski studij znanosti o okolišu (Undergraduate Study of Environmental Sciences) Mentor: prof.dr.sc. Zlatko Liber Zagreb, 2010. SADRŽAJ 1. UVOD........................................................................................2 2. Porodica Droseraceae.................................................................3 2.1. Rod Dionaea L..................................................................3 2.2. Rod Drosera L...................................................................5 2.3. Rod Aldrovanda L.............................................................7 3. Porodica Sarraceniaceae.............................................................8 3.1. Rod Darlingtonia L............................................................8 3.2. Rod Heliamphora L............................................................9 3.3. Rod Sarracenia L..............................................................10 4. Porodica Nepenthaceae...............................................................11 4.1. Rod Nepenthes L................................................................11 5. Porodica Lentibulariaceae..........................................................12 5.1. Rod Genlisea L..................................................................12 5.2. Rod Pinguicula L..............................................................13
    [Show full text]
  • Phylogeny and Biogeography of South American Marsh Pitcher Plant Genus Heliamphora (Sarraceniaceae) Endemic to the Guiana Highlands
    bioRxiv preprint doi: https://doi.org/10.1101/2020.04.29.068395; this version posted April 30, 2020. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC-ND 4.0 International license. Phylogeny and Biogeography of South American Marsh Pitcher Plant Genus Heliamphora (Sarraceniaceae) Endemic to the Guiana Highlands Sukuan Liu and Stacey D. Smith Author for correspondence: Sukuan Liu, [email protected] Department of Ecology and Evolutionary Biology, University of Colorado Boulder, 1900 Pleasant Street, Boulder, Colorado 80309, U.S.A. Abstract: Heliamphora is a genus of carnivorous pitcher plants endemic to the Guiana Highlands with fragmented distributions. We presented a well resolved, time-calibrated, and nearly comprehensive Heliamphora phylogeny estimated using Bayesian inference and maximum likelihood based on nuclear genes (26S, ITS, and PHYC) and secondary calibration. We used stochastic mapping to infer ancestral states of morphological characters and ecological traits. Our ancestral state estimations revealed that the pitcher drainage structures characteristic of the genus transformed from a hole to a slit in single clade, while other features (scape pubescence and hammock-like growth) have been gained and lost multiple times. Habitat was similarly labile in Heliamphora, with multiple transitions from the ancestral highland habitats into the lowlands. Using Mantel test, we found closely related species tend to be geographically closely distributed. Placing our phylogeny in a historical context, major clades likely emerged through both vicariance and dispersal during Miocene with more recent diversification driven by vertical displacement during the Pleistocene glacial-interglacial thermal oscillations.
    [Show full text]
  • Photosynthetic Characterization of Australian Pitcher Plant Cephalotus Follicularis
    DOI: 10.1007/s11099-011-0032-0 PHOTOSYNTHETICA 49 (2): 253-258, 2011 Photosynthetic characterization of Australian pitcher plant Cephalotus follicularis A. PAVLOVIČ Department of Plant Physiology, Faculty of Natural Sciences, Comenius University in Bratislava, Mlynská dolina B2, 842 15, Bratislava, Slovakia Abstract Australian carnivorous pitcher plant Cephalotus follicularis Labill. produces two types of leaves. During the spring time, the plant produces a foliage type of noncarnivorous leaf called lamina. Later, the second type of leaf is produced - carnivorous pitcher. Using simultaneous measurements of gas exchange and chlorophyll (Chl) fluorescence photosynthetic efficiency of these two distinct forms of leaves were compared. In addition stomatal density, an important component of gas exchange, and Chl concentration were also determined. Pitcher trap had lower net photosynthetic rate (PN) in comparison to noncarnivorous lamina, whereas the rate of respiration (RD) was not significantly different. This was in accordance with lower stomatal density and Chl concentration in the pitcher trap. On the other hand maximum quantum yield of PSII (Fv/Fm) and effective quantum yield of photochemical energy conversion in PSII (ΦPSII) was not significantly different. Nonphotochemical quenching (NPQ) was significantly higher in the lamina at higher irradiance. These data are in accordance with hypothesis that changing the leaf shape in carnivorous plants to make it a better trap generally makes it less efficient at photosynthesis. However, the pitcher of Cephalotus had much higher PN than it was expected from the data set of the genus Nepenthes. Because it is not possible to optimize for contrasting function such as photosynthesis and carnivory, it is hypothesized that Cephalotus pitchers are less elaborated for carnivorous function than the pitchers of Nepenthes.
    [Show full text]
  • Carniflora News – February 2020 (PDF)
    THE AUSTRALASIAN CARNIVOROUS PLANTS SOCIETY INC. CARNIFLORA NEWS A.B.N. 65 467 893 226 February 2020 Aldrovanda vesiculosa in flower. Photographed by David Colbourn Drosera petiolaris. Photographed by Robert Gibson Welcome to Carniflora News, a newsletter produced by the Australasian Carnivorous CALENDAR Plants Society Inc. that documents the meetings, news and events of the Society. FEBRUARY The current committee of the Australasian Carnivorous Plant Society Inc. comprises: 7th February 2020 - AUSCPS meeting - Canberra featuring a Venus Fly Trap workshop 9th February 2020 - Old Bus Depot Markets - Canberra 14th February 2020 - AUSCPS meeting - Sydney Utricularia, Aldrovanda & Genlisea COMMITTEE MARCH 6th March 2020 - AUSCPS meeting - Canberra featuring Carnivorous Plants 101 President - Wesley Fairhall 13th March 2020 - AUSCPS meeting - Sydney Byblis, Drosophyllum & Roridula [email protected] 15th March 2020 - Old Bus Depot Markets - Canberra 28-29th March - Collectors’ Plant Fair, Clarendon, N.S.W. Vice President - Barry Bradshaw APRIL [email protected] 3rd April 2020 - AUSCPS meeting - Canberra Utricularia, Aldrovanda & Genlisea 10th April 2020 - AUSCPS meeting - Sydney Nepenthes & carnivorous bromeliads Treasurer - David Colbourn 13th April 2020 - Royal Easter Show - Carnivorous Plant Competition [email protected] MAY 1st May 2020 - AUSCPS meeting - Canberra Cephalotus, Heliamphora and Pinguicula Secretary - Kirk ‘Füzzy’ Hirsch 8th May 2020 - AUSCPS meeting - Sydney featuring Cephalotus and Heliamphora [email protected] JUNE General Committee Member - Sean Polivnick 5th June 2020 - AUSCPS meeting - Canberra Pygmy Drosera, perennial Byblis & [email protected] Roridula 12th June 2020 - AUSCPS meeting - Sydney featuring Carnivorous bromeliads JULY 3rd July 2020 - AUSCPS meeting - Canberra featuring a Sarracenia and Darlingtonia DELEGATES 10th July 2020 - AUSCPS meeting - Sydney (AGM) featuring Winter growing Drosera AUGUST Journal Editor - Dr.
    [Show full text]
  • The Roots of Carnivorous Plants
    Plant and Soil (2005) 274:127–140 Ó Springer 2005 DOI 10.007/s11104-004-2754-2 The roots of carnivorous plants Wolfram Adlassnig1, Marianne Peroutka1, Hans Lambers2 & Irene K. Lichtscheidl1,3 1Institute of Ecology and Conservation Biology, University of Vienna, Althanstrasse 14, 1090 Vienna, Austria. 2School of Plant Biology, Faculty of Natural and Agricultural Sciences, The University of Western Australia, Crawley WA 6009, Australia. 3Corresponding author* Received 30 April 2004. Accepted in revised form 31 August 2004 Key words: carnivorous plants, insectivorous plants, morphology, nutrition, root Abstract Carnivorous plants may benefit from animal-derived nutrients to supplement minerals from the soil. Therefore, the role and importance of their roots is a matter of debate. Aquatic carnivorous species lack roots completely, and many hygrophytic and epiphytic carnivorous species only have a weakly devel- oped root system. In xerophytes, however, large, extended and/or deep-reaching roots and sub-soil shoots develop. Roots develop also in carnivorous plants in other habitats that are hostile, due to flood- ing, salinity or heavy metal occurance. Information about the structure and functioning of roots of car- nivorous plants is limited, but this knowledge is essential for a sound understanding of the plants’ physiology and ecology. Here we compile and summarise available information on: (1) The morphology of the roots. (2) The root functions that are taken over by stems and leaves in species without roots or with poorly developed root systems; anchoring and storage occur by specialized chlorophyll-less stems; water and nutrients are taken up by the trap leaves. (3) The contribution of the roots to the nutrient supply of the plants; this varies considerably amongst the few investigated species.
    [Show full text]
  • SOME PERSONAL OBSERVATIONS on CULTIVATING the HELIAMPHORA by Robert R
    SOME PERSONAL OBSERVATIONS ON CULTIVATING THE HELIAMPHORA by Robert R. Ziemer (P. O. Box 4562, Arcata, CA 95521) The following note is based on some 7 plants easily topple. I have tried clay years experience growing three species of pots, but, for some reason, the plants in Heliamphora - H. heterodoxa. H. nu- plastic pots have grown much better. I tans, and H. minor. This information is have also tried 25 x 40 cm plastic dish not intended to be a definitive or even a tubs with the bottom filled with perlite comprehensive guide to the cultivation of and the top 3/4 with live sphagnum. Again, these species, but simply some observa- the plants in the 20 cm pots grow much tions on what I have found to work for me better. through trial and error. I have not My Heliamphora absolutely do not conducted any rigorously controlled ex- grow well in water-logged conditions. periments on either potting mixture, They also do not like to be subjected to light, or temperature. Since these plants standing water. Drainage is a must. One are sufficiently difficult to obtain, every problem I have found with live sphag- death is a traumatic experience for me num is that even though the surface is and I have not wanted to repeat the se- green and growing, a few cm below the quence simply to study the reasons for surface the moss begins to break down failure. My main objective has been to and, within a year, forms a rather water- keep the little fellows alive rather than logged medium.
    [Show full text]
  • Carnivorous Plantsplants –– Classicclassic Perspectivesperspectives Andand Newnew Researchresearch
    CarnivorousCarnivorous plantsplants –– classicclassic perspectivesperspectives andand newnew researchresearch Barry Rice The Nature Conservancy, Davis, USA The ranks of known carnivorous plants have grown to approximately 600 species. We are learning that the relationships between these feeders and their prey are more complex, and perhaps gentler, than previously suspected. Unfortunately, these extraordinary life forms are becoming extinct before we can even document them! Carnivorous plants are able to do four things: they attract, false signals, the trigger hairs must be bent, not once, but trap, digest and absorb animal life forms. While these four two or more times in rapid succession. In effect, the plant abilities may seem remarkable in combination, they are, can count! When the trap first closes, the lobes fit together individually, quite common in the plant kingdom. All very loosely, the marginal spines interweaving to form a plants that produce flowers for the purpose of summoning botanical jail. Prey items that are too small to be worth pollinators are already skilled at attracting animals. Many digesting can quickly escape, and the trap will reopen the plants trap animals at least temporarily, usually for the next day. But, large prey remain trapped, and their purposes of pollination. Digestion may seem odd, but all panicked motions continue to stimulate the trigger hairs. plants produce enzymes that have digestive capabilities – This encourages the traps to seal completely, suffocating carnivorous plants have only relocated the site of enzy- the prey, and to release digestive enzymes. (Children who matic activity to some external pitcher or leaf surface. feed dead flies to their pet Venus flytraps are often disap- Finally, absorption of nutrients is something that all pointed when, the next day, the uninterested plants open plants do (or, at least, all that survive past the cotyledon their traps and reject the inanimate morsels – only live stage).
    [Show full text]
  • Carnivorous Plant Newsletter V46 N4, December 2017
    New cultivars Keywords: cultivar, Dionaea muscipula ‘Red Pico-Teeth’, Heliamphora ‘Patasola’, Nepenthes ampullaria ‘Black Miracle’, Nepenthes ampullaria ‘Black Pearl’. Dionaea muscipula ‘Red Pico-Teeth’ Submitted: 16 July 2017 Dionaea muscipula ‘Red Pico-Teeth’ was obtained as a seedling from Lucien Blacher in De- cember 2013. The entire plant is dark red and the leaves are prostrate rather than erect (Fig. 1). The traps have extremely short, even non-existent teeth — as opposed to D. ‘Red Micro-Teeth’ and other cultivars that have larger teeth. The name is derived from this cultivar being all red and pico (for picometer) refers to the traps having extremely short or non-existent teeth. This plant should be reproduced only by vegetative means to ensure that its unique characteris- tics are maintained. Figure 1: Dionaea muscipula ‘Red Pico-Teeth’ plant and trap. —Sébastien Bonnet • 20 hameau des noëls • 10000 TROYES • France • [email protected] Heliamphora ‘Patasola’ Submitted: 18 June 2017 Heliamphora ‘Patasola’ (Fig. 2 and Back Cover) is a hybrid H. parva × folliculata that I created several years ago. It is unique among its siblings based on its vigor, rapid growth rate, and unique physical characteristics. It grew from seed to flowering adult in under four years. Jennifer Lei culti- vated this particular seedling to maturity. Mature pitchers are 25-30 cm tall and 5-6 cm wide. Under intense lighting, the pitcher color will become vinaceous to violet to almost charcoal as the pitcher ages. Pitcher shape is infundibular in the lower part with a pronounced waist approximately half way up. The upper section is infundibular and slightly compressed in older pitchers from front to back.
    [Show full text]